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DOI: 10.1038/ncomms12306
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Protected areas are widely considered essential for biodiversity conservation. However, few
global studies have demonstrated that protection benets a broad range of species. Here,
using a new global biodiversity database with unprecedented geographic and taxonomic
coverage, we compare four biodiversity measures at sites sampled in multiple land uses
inside and outside protected areas. Globally, species richness is 10.6% higher and abundance
14.5% higher in samples taken inside protected areas compared with samples taken outside,
but neither rarefaction-based richness nor endemicity differ signicantly. Importantly, we
show that the positive effects of protection are mostly attributable to differences in land use
between protected and unprotected sites. Nonetheless, even within some human-dominated
land uses, species richness and abundance are higher in protected sites. Our results reinforce
the global importance of protected areas but suggest that protection does not consistently
benet species with small ranges or increase the variety of ecological niches.
1 School of Life Sciences, University of Sussex, Falmer, Brighton BN1 9QG, UK. 2 United Nations Environment Programme World Conservation Monitoring
Centre, 219 Huntingdon Road, Cambridge CB3 0DL, UK. 3 Department of Life Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, UK.
4 Department of Biosciences, College of Science, Swansea University, Singleton Park, Swansea SA2 8PP, UK. 5 CSIRO Land and Water, Canberra Australian
Capital Territory 2601, Australia. 6 Department of Life Sciences, Imperial College, London, Silwood Park, London SL5 7PY, UK. * These authors contributed
equally to this work. w Present address(es): Conservation Programmes, Zoological Society of London, London NW1 4RY, UK (C.L.G.); Centre for Biodiversity
and Environment Research, Department of Genetics, Evolution and Environment, University College London, Gower Street, London WC1E 6BT, UK (T.N.).
Correspondence and requests for materials should be addressed to C.L.G. (email: claudiagray@gmail.com) or to S.L.L.H. (email: samantha.hill@nhm.ac.uk) or
to J.P.W.S. (email: j.scharlemann@sussex.ac.uk).
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23.5
23.5
All sites
All sites
Matched sites
Matched sites
75
Latitude
50
25
0
25
1,500
1,000
500
500
1,000
1,500
50
Number of sites
Protected
Management
Category
Group
Unprotected 4,397
All protected 1,781
631
III - VI
Unknown
746
I & II
404
Species richness
difference (%)
10 0
Rarefied richness
difference (%)
10 20 30 40
**
*
*
*
10 0
3,761
1,627
570
623
434
10 20 30 40
10 0
*
165
2,448
859
352
378
129
10 20 30 40
Endemicity
difference (%)
10 0
10 20 30 40
4,114
1,744
628
768
348
Figure 2 | Effects of terrestrial protected areas on four local biodiversity measures. (a) Species richness, (b) total abundance, (c) rareed richness and
(d) endemicity at sites inside (lled circles) relative to sites outside protected areas (open). Estimates are given separately for protected areas in different
management category groups (grey circles; least restrictive (IUCN categories IIIVI), unknown (missing IUCN category, potentially mixed set of categories)
and most restrictive land management regimes (IUCN categories I and II)). Bars indicate 95% CIs; **Po0.01 and *Po0.05. Number of sampled sites in each
category is shown; sample sizes vary between panels due to differences in the use of occurrence and abundance data to calculate biodiversity measures (see
Supplementary Information). Separate generalized linear mixed effects models were run for each response variable (see Supplementary Information for further
information). Supplementary Fig. 3 gives the corresponding results for analyses where sites were matched across the protected area boundary by land use.
NATURE COMMUNICATIONS | 7:12306 | DOI: 10.1038/ncomms12306 | www.nature.com/naturecommunications
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20
760
238
613
115
276
33
530
279
26
666
297
415
132
80
261
740
236
186
67
20
1,220
819
40
40
40
20
20
99
599
213
344
65
20
12
636
203
584
108
249
31
606
232
388
137
213
75
40
68
31
937
719
40
20
20
442
122
289
62
413
146
43
18
173
46
40
256
78
538
321
40
20
20
Primary
Secondary
579
283
171
58
40
1,110
759
Plantation Cropland
Pasture
Urban
Human dominatad
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Species richness
difference (%)
Unprotected
Protected
20
10
0
10
3479
372
191
326
322
Unprotected
Small,
young
Small,
old
Large,
young
Large,
old
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outside protected areas were caused by biases in the location of protected areas27,
we considered elevation47 and derived slope at c. 1 km2 resolution and agricultural
suitability48 at 10 km2 resolution as covariates in all models (see Supplementary
Information for further details). To ensure independence of all variables
in the model, we intentionally included only these three confounding variables that
we considered to be fully independent of the presence of a protected area.
For example, distances to roads and markets are affected by the presence of
protected areas so are not independent confounding factors (see Supplementary
Information for details). We sequentially compared models with and without each
xed effect and at each step dropped the term with the highest P-value, until all
terms had Po0.05 (ref. 49).
Assessing protection effects. We tested for biodiversity differences between sites
inside and outside protected areas using the all-sites data, treating protection status
(inside vs outside a protected area) as a xed effect. We then tested whether
biodiversity measures differed between management category groups by re-coding
IUCN category as a four-level factor: unprotected, IUCN category IIIVI, IUCN
category unknown, and IUCN category I and II.
Assessing protection effects within and among land uses. We used two
approaches to test whether biodiversity differences between protected and
unprotected sites varied with land use. First, using the all-sites data, we modelled
the response of each biodiversity measure to protection status, land use, and their
interaction. We also tested for the three-way interaction between land use,
protection and either use intensity, latitudinal zone or taxonomic group. Second,
using the matched-sites data, we re-ran models with protection status, and then
with management category group as a xed effect. We also split the matched-sites
data by latitudinal zone and taxonomic groups to assess whether these factors
inuenced the effect of protection. Finally, we tested whether the site-level
biodiversity response to protection varied with the size/age class of the protected
area [four-level factor with all combinations of young (o20 years), old (2085
years), small (o400 km2) and large (40012,000 km2); these thresholds between
categories were selected to give a similar number of sites in each group].
Estimating global protected area effectiveness. The global effectiveness of
protected areas (e) was estimated from e 1 (1 i)/(1 o), where modelled
site-level biodiversity inside (i) and outside (o) protected areas are expressed as a
proportion of that under pristine conditions. We calculated the ratio of i/o from
the model estimates for biodiversity inside relative to outside protected areas in
each land use (Fig. 3), where each land-use parameter was weighted by the
proportion of global terrestrial area within that land-use type. This value of i/o
could then be used to solve an equation expressing the global state of site-level
biodiversity: 1 r ai (1 a)o, where r is the estimated global average loss of
site-level biodiversity relative to pristine46 and a is the fraction of the total land area
that is protected50. Solving this equation for i and o allowed us to estimate e.
Finally, by using estimates for the effect of protection in IUCN categories I and II
(Fig. 2a,b) to give i/o, we estimated e under the more restrictive management
scenario. By rearranging the equations we estimated the total protected area (a)
needed to obtain the same average local biodiversity outcome (1 r) inferred
under this more restrictive management scenario. See Supplementary Information
for more details.
Data availability. The biodiversity data that support the ndings of this study
are available in the Natural History Museum data portal (data.nhm.ac.uk)
with the identier dx.doi.org/10.5519/0095544. R scripts are available at
http://github.com/claudialouisegray/PREDICTS_WDPA.
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Acknowledgements
We thank the hundreds of data contributors; all PREDICTS project volunteers, masters
and PhD students that collated records; and Adriana De Palma, Helen Phillips, Diego
Juffe-Bignoli, Neil Burgess, Max Gray, Daniel Ingram, Valerie Kapos, Naomi Kingston,
Sarah Luke and the protected areas team at UNEP-WCMC for comments and assistance.
We thank the School of Life Sciences at the University of Sussex for support and the
Natural History Museum for a GIA travel award. The PREDICTS project is funded by
the UK Natural Environment Research Council (NERC, grant number: NE/J011193/2).
PREDICTS is endorsed by the Group on Earth Observations Biodiversity Observation
Network (GEO BON). This is a contribution from the Imperial College Grand
Challenges in Ecosystem and the Environment Initiative, and the Sussex Sustainability
Research Programme.
Author contributions
C.L.G., S.L.L.H., T.N., A.P. and J.P.W.S. designed research; C.L.G., S.L.L.H., T.N., L.B.,
A.P. and J.P.W.S. performed research; C.L.G., S.L.L.H., T.N., L.N.H., S.C., A.J.H., S.F.,
A.P. and J.P.W.S. contributed data and analytical tools; C.L.G. and S.L.L.H. analysed data;
C.L.G., S.L.L.H., T.N., L.N.H., L.B., A.P. and J.P.W.S. wrote the paper.
Additional information
Supplementary Information accompanies this paper at http://www.nature.com/
naturecommunications
Competing nancial interests: The authors declare no competing nancial interests.
Reprints and permission information is available online at http://npg.nature.com/
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How to cite this article: Gray, C. L. et al. Local biodiversity is higher inside than outside
terrestrial protected areas worldwide. Nat. Commun. 7:12306 doi: 10.1038/ncomms12306
(2016).
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